In southwestern England, red deer, Cervus elaphus, are culled by rifle (‘stalking’) or by hunting with hounds (‘hunting’). We compare the welfare costs of the two culling methods. Observations of hunts revealed that likely stressors such as close proximity to humans and hounds, active pursuit, noise, obstruction and physical restraint prior to despatch were very common. Other stressors, such as wounding, were rare. The blood profiles of hunted deer were compared both with injured deer, which were put down because they were thought to be suffering, and with stags stalked in the rutting season, when mature males rapidly lose weight and may be damaged in fights. Extensively hunted deer did not differ from severely injured deer in measures of muscle disruption: in hunted deer measures of red blood cell damage and psychological stress were higher. Hunted stags killed during the rut showed markedly higher levels of measures of blood and muscle cell disruption, psychological stress and fat reserve mobilization than stalked stags killed during this season. Estimates of wounding rates by stalkers showed that 11 per cent of deer required two or more shots to kill, 7 per cent took 2—15 min to die and 2 per cent escaped wounded. Overall, we judged that the welfare costs associated with hunting red deer were higher than those associated with stalking, and reducing the welfare costs associated with hunting was much less feasible than reducing those associated with stalking.

Our study aimed at examining the phylogenetic position of the newly-found Setaria nematodes obtained from the red deer (Cervus elaphus) based on sequences of the mitochondrial cytochrome c oxidase subunit 1 (COX-1). Alignment and phylogenetic analyses, as well as SEM microscopic analysis, revealed the presence of two Setaria species: S. cervi and S. tundra. Setaria tundra was noted in only one individual, a calf of the red deer, while S. cervi was observed in three stages, two hinds and one calf of the red deer. According to our knowledge, it is the first case of S. cervi in the red deer in Poland confirmed in molecular studies and also the first case of S. tundra infection in the red deer.

The main factors affecting the mechanical (and other) properties of bone, including antler, are the proportions of ash (especially Ca and P) and collagen content. However, some trace minerals may also play more important roles than would be expected, given their low levels in bone and antler. One such trace mineral is Cu. Here, we studied the effects of Cu supplementation on the mechanical and structural characteristics, and mineral content of antlers from yearling and adult (4 years of age) red deer fed a balanced diet. Deer (n=35) of different ages (21 yearlings and 14 adults) were studied. A total of 18 stags (11 yearlings and 7 adults) were injected with Cu (0.83 mg Cu/kg BW) every 42 days, whereas the remaining 17 (10 yearlings and 7 adults) were injected with physiological saline solution (control group). The Cu content of serum was analysed at the beginning of the trial and 84 days after the first injection to assess whether the injected Cu was mobilized in blood. Also, the mechanical and structural properties of antlers and the mineral content in their cortical walls were examined at three (yearlings) or four (adults) points along the antler beam. The effect of Cu supplementation was different in yearlings and adults. In yearlings, supplementation increased the Cu content of serum by 28%, but did not affect antler properties. However, in adults, Cu supplementation increased the Cu content of serum by 38% and tended to increase the cortical thickness of antlers (P=0.06). Therefore, we conclude that, even in animals receiving balanced diets, supplementation with Cu could increase antler cortical thickness in adult deer, although not in yearlings. This may improve the trophy value of antlers, as well as having potential implications for bones in elderly humans, should Cu supplementation have similar effects on bones as those observed in antlers.

Deer exclosure is an important management strategy for encouraging woodland regeneration in the presence of high numbers of Cervus elaphus L. This could pose a threat to important saxicolous lichen communities as a result of competition from bryophytes and other vegetation. This investigation compared the bryophyte and lichen communities associated with siliceous rock outcrops and boulders inside and outside a number of exclosures in wet heath vegetation at the Gruinard, Letterewe and Little Gruinard estates in Wester Ross. Species cover data were recorded from 6600·2×0·2 m quadrats, allocated to three different aspects of rocks, from 22 pairs of plots (placed either side of a deer fence) with randomization at each level of this hierarchy. The data were analyzed using linear mixed effects models. Mean lichen cover, diversity and species richness were significantly lower in quadrats on rocks within the exclosures. Mean bryophyte cover was significantly higher in quadrats on rocks inside the deer fence and was shown to have a highly significant negative relationship with total lichen cover. Saxicolous lichen species are particularly affected by the increase in shading and litter accumulation from the surrounding plants and the subsequent growth of bryophytes and heath vegetation on rocks within the exclosures. The results of this study have implications for the use of exclosures at locations where there are important saxicolous lichen communities.

The relationships among harem size, adult sex ratio (proportion of males >5 years in the adult population, i.e. males >5 years plus females >2 years) and male age-structure of red deer Cervus elaphus were investigated in La Petite Pierre National Reserve (PPNR) in France. We tested whether: (1) increasing adult sex ratio leads to a decrease in harem size along with an increase in the number of harems within a given rut period; (2) whether participation of sub-adult males in mating activities increases with decreasing adult sex ratio, and as the proportion of adult males decreases. Harem size did not vary over the mating period, suggesting a high turnover of harem-holders leading to an increase in the costs of mating for males. Harem size averaged 1.43±0.91 and was lower than harem sizes typically reported for red deer (e.g. >2.5 in Scotland and Norway). In support of the first prediction, a decrease in harem size and an increase in the total number of harems seen with an increasing sex ratio was observed (harem size=2.08−1.26 [±0.43]×(sex ratio); r2=0.25, F1,18=6.19, P=0.02). Both the uniform distribution of females among harem stags and the small harem sizes observed in PPNR might concur to a smaller variance in male reproductive success than previously reported in red deer. Lastly, in partial support of the second prediction, the proportion of sub-adult males observed during the mating season decreased with increasing adult sex ratio and with increasing proportions of adult males. Whether or not the lower proportion of sub-adults seen when competition among mature males increases means that fewer young males mate cannot be assessed from our study.

In ungulates, a set of deciduous teeth is replaced by a set of permanent teeth during their first 2 years of life. Little is known about factors causing variation in timing of tooth eruption, but overall size, condition or mineral deficiencies may play a role. The pattern of incisor (I) and canine (C) eruption was investigated in 2241 yearlings from five populations of red deer Cervus elaphus in Norway during 1964–2001. Before the sampling period (10 September–15 November), I1 had erupted in most yearlings (97%), while eruption of I3 and C had barely started (erupted in 7% and 5% of individuals, respectively). Exactly 50% of the yearlings had erupted I2. Jawbone length was shown to be a main determining factor for eruption of I2. However, for a given jawbone length, there was large variation among populations in the proportion of yearlings with erupted I2 (from <20% to >75%). Females got their permanent I2 earlier than males. Tooth eruption was found to be later with increasing local deer density, even after correcting for body weight. There was a residual effect of body weight (a measure of condition) after body size (measured as jawbone length) was controlled for in one of five populations, indicating that overall condition may influence timing of tooth eruption in this area. There was no evidence that calcium was limiting for eruption, as no correlation was found between timing of eruption and limestone in bedrock. Since there is large variation in timing of tooth eruption among red deer populations even after controlling for differences in phenotype and environmental factors, genotypic variation may also play an important role for tooth eruption patterns.

In Mediterranean environments browse is expected to be a major component of the diet of red deer Cervus elaphus during the summer when most grasses senesce. In this study the hypothesis is tested that, if browse is an essential food resource at this time, the sex with the larger body size may have a greater physical ability to reach the tree canopy and include a higher proportion of browse in its diet. The organic matter digestibility (OMD) and composition of the diet of a population of red deer males and females were measured between July and September, in 1996 and 1997, in an area of Portugal with a Mediterranean type of climate, using the n-alkane technique. Direct observations of animals browsing on tree canopies and shrubs were made also during the same period. The OMD of the diet was significantly different between sexes and between years and was higher in males than females both in 1996 (means ± sem for males and females, 0.65 and 0.58 ± 0.026, respectively) and 1997 (means for males and females, 0.59 and 0.48 ± 0.045, respectively). The proportion of cork oak Quercus suber in the diet was also significantly different between sexes and between years, with males having a higher proportion of cork oak in their diets in 1996 (males and females, 0.30 and 0.16 ± 0.037, respectively) and 1997 (males and females, 0.41 and 0.23 ± 0.064, respectively). A significantly lower proportion of the herbage layer was found in the diet of males in 1996 (males and females, 0.46 and 0.64 ± 0.048) but not in 1997. The number of observations of males browsing on tree canopies was significantly higher than females. It is concluded that there are differences between sexes in the composition and quality of the diet and that a larger body size may confer advantages in situations where browse is an essential food resource.

Inter-sexual segregation between habitats or between sites within a habitat is very widespread in sexually dimorphic ungulates. As an explanation, it has been suggested that males, because of their larger forage requirements, are driven from preferred, high quality forage habitats into marginal habitats of lower forage quality but higher forage biomass by female competition (‘indirect competition hypothesis’). However, this hypothesis has hardly been tested. In the present study, we tested the indirect competition hypothesis for site segregation of red deer on the Isle of Rum, Scotland, within two vegetation communities: short Agrostis/Festuca grassland and Calluna heath. We predicted that: (1) males should choose sites within each vegetation community that are of higher forage biomass but lower forage quality, than sites selected by females; (2) the degree of inter-sexual site segregation should be highest within the vegetation community in which indirect competition was most intense. We found that the sexes were significantly segregated between sites within both vegetation communities, but males did not use sites of lower forage quality or of higher forage biomass than did females. Moreover, degree of site segregation was not significantly higher in the vegetation community in which competition was more intense (i.e. short Agrostis/Festuca grassland). We conclude that the indirect competition hypothesis cannot explain site segregation in red deer on the Isle of Rum.

Three groups of factors which might affect the blood biochemistry of red deer (Cervus elaphus) were examined. These were the wounding site (head/neck versus chest), the stalker who collected the blood (coupled with the geographical area where each deer was shot), and the sex and nutritional status of the deer. The activities of muscle enzymes, creatine kinase, aspartate amino transferase, and lactate dehydrogenase, were markedly higher in the plasma of deer shot in the chest as compared with those shot in the head or neck; the plasma also looked different, being deep cherry red in colour in the chest-shot deer. Other biochemical measures were unaffected by the wounding site. Blood collected by one stalker from the chest cavity had higher activity of the liver enzyme, glutamate dehydrogenase, higher concentration of potassium and lower concentrations of sodium and chloride than blood collected by another stalker from a knife wound in the base of the neck. Otherwise the method of collecting blood had very little effect on blood biochemistry. Cortisol was unaffected by the wound site nor, probably, by the method of collecting blood, but was highest in the stags living in a more mountainous region and in those deer that had to be shot twice before they died. Stags, which were shot in the rutting season and were probably fasting, had significantly higher concentrations of free fatty acids than hinds. Lactating hinds had significantly less fat on their kidneys than non-lactating hinds and stags. All hinds, which were shot in winter, had lower concentrations of urea than stags, which were shot in the autumn.

The use of a seaweed habitat by red deer Cervus elaphus L. on the Isle of Rum, Scotland, was examined in detail. New information is provided on diet selection, timing of seaweed use relative to tides, inter-individual differences in seaweed use, and sex differences in site use within the seaweed habitat (‘site segregation’). Interestingly, seaweed use by adult males and females was closely correlated to that of their mothers. This implies that deer ‘learn’ early in life to include seaweed into their diet. Formerly, it has been suggested that male inferiority in indirect competitive ability relative to females causes site segregation in dimorphic ungulates (‘indirect competition hypothesis’). The observed pattern of site segregation within the seaweed habitat was used to test the hypothesis, which predicts that males should be found at sites where they can achieve higher intake rates, but where forage quality is lower than at female sites. With respect to seaweed use, the hypothesis further predicts that segregation should be lower within the seaweed habitat than within terrestrial vegetation communities, and that males should time seaweed use earlier (relative to the tide) than females. This is because seaweed availability is more subject to tidal rhythm than to indirect competition in comparison to terrestrial habitats. Males and females used different bays, and within bays they used different fractions of seaweed. However, male-preferred sites did not yield higher intake rates and were not of lower forage quality than sites preferred by females. Moreover, segregation was not lower within the seaweed habitat than within terrestrial vegetation communities, and males did not time their seaweed use earlier relative to the tide than did females. The indirect competition hypothesis could not explain the observed pattern of site segregation. Other factors, such as sex differences in sheltering or anti-predator behaviour, or social harassment, could be responsible instead.

Rutting calls of male and alarm calls of female red deer (Cervus elaphus), sika deer (C. nippon), and their hybrids were recorded from wild and captive-bred populations. Vocal variations between these species and their hybrids (C. elaphus × C. nippon) were examined using multivariate analysis to determine whether these groups could be successfully discriminated. There were highly significant differences between the vocalizations of each group for both sexes (males: Wilks' Λ = 0.0233, P<0.0001; females: Wilks' Λ = 0.1338, P<0.0001). The greatest differences were between analogous calls of red and sika deer. Vocalizations of hybrids from both captive-bred and feral populations were intermediate between the two parent species. The differences between the groups were most highly influenced by the pitch of calls. The results of this study indicate that the vocalizations of deer, rather than being learned or mimicked from similar species in their proximity, are strongly influenced by the genetic background of their parents.

Intake, rumen digesta pool sizes, ruminal digestion rates, passage rates of liquid and particle pools and faecal particle-size distrubutions of grass, browse and lucerne diets were compared in ruminally fistulated moose, wapiti and cattle. For each diet, ruminal digestion rates (nylon bag technique) were similar for the three ruminants. Intakes, rumen digesta pool sizes and passage rates varied among species and higher rates were associated with passage of larger faecal particles. However, it was not possible to simply rank the three ruminants as representative grazers (cattle), browsers (moose) and mixed feeders (wapiti) since intakes, passage rates and rumen pool sizes interacted with diet and season. Contrary to expectation, digestible dry matter intakes of each species were not greatest for their respective typical diets.